Soil organic carbon, total nitrogen stocks and CO2 emissions in top- and subsoils with contrasting management regimes in semi-arid environments

This study aims to investigate soil organic carbon (SOC) and total nitrogen (TN) contents and stocks, CO2 emissions and selected soil properties in croplands, grazing lands, exclosures and forest lands of semi-arid Ethiopia. Sampling was done at 0–30, 30–60 and 60–90 cm soil depths and concentration and stocks of SOC, TN and selected soil properties were determined using standard routine laboratory procedures. There were variations in distribution of SOC and TN stock over 90 cm depth across land use types and locations, decreasing from topsoils to subsoil, with average values ranging from 48.68 Mg C ha−1 and 4.80 Mg N ha−1 in Hugumburda cropland to 303.53 Mg C ha−1 and 24.99 Mg N ha−1 in Desa’a forest respectively. Forest sequestered significant higher SOC and TN stock, decreasing with depth, compared with other land use types. In Desa'a and Hugumburda, the conversion of forest to cropland resulted in a total loss of SOC stock of 9.04 Mg C ha−1 and 2.05 Mg C ha−1, respectively, and an increase in CO2 emission of 33.16 Mg C ha−1 and 7.52 Mg C ha−1 yr−1, respectively. The establishment of 10 years (Geregera) and 6 years (Haikihelet) exclosures on degraded grazing land increased SOC stock by 13% and 37% respectively.

(i) To quantify the distribution of SOC and TN concentrations and stocks and CO 2 emissions within the profile to a depth of 90 cm in 30 cm increments in of four contrasting land use types (forest, exclosure, grazing land and cropland); (ii) To quantify variations in selected soil properties (total nitrogen, pH, cation exchange capacity (CEC), particle size distribution and bulk density) in these land use types; and. (iii) To identify biophysical controls (vegetation, soil properties, climate) on depth distribution of SOC, N storage and CO 2 emission in tropical semi-arid soils of northern Ethiopia.
We hypothesized that (i) the impact of vegetation cover/land use on SOC and TN stocks and CO 2 emissions in these soils is not limited to the topsoil layer, (ii) climate and land use will serve as possible indicators to modulate the magnitude of changes in SOC and TN stocks, (iii) SOC and TN concentrations and stocks decrease following conversion of forest to grazing land and cropland, but increase with establishment of exclosure on degraded grazing land.

Materials and methods
Description of the study area. Four locations ( Fig. 1) with different land use types were chosen for this study: Hugumburda (natural forest, grazing land and cropland, 2494 m.a.s.l), Desa'a (natural forest, grazing land and cropland, 2433 m.a.s.l), Geregera watershed (exclosures, grazing land and cropland, 2180 m.a.s.l) and Haikihelet watershed (exclosures, grazing land and cropland, 2236 m.a.s.l), all in the semi-arid area of Tigray Regional State, northern Ethiopia ( Fig. 2; Table 1). Exclosure refers to previously degraded grazing land, which has been fully conserved without any form of animal and human activity as a sustainable way of restoring land degradation through natural regeneration 41 . The age durations of the exclosures in the Haikihelet and Geregera watersheds are 6 and 10 years respectively 42 . Cambisols are the dominant soil types in Hugumburda, Haikihelet and Geregera, while Vertisols dominate in Desa'a 43 .
In the study area, the mean annual precipitation (MAP) between 1983 and 2016 is about 503 mm 44 . The rainy season peak period is in July/August and rescinds towards September. The estimated average temperature in the region is 18 °C, with significant variations with altitude. The study sites were classified as mid-altitude (1800-2200 m above sea level) and high altitude (> 2100 m above sea level) classes, based on the traditional indigenous agro-climate classification system in Ethiopia.
The most common crop rotation in the study area is wheat + barley + faba bean/field pea, while teff + maize can be switched after a few years. At the time of soil sampling, all cropland was under rainfed cereal crop cultivation. Sampling took place between November and December after harvest, when the soil conditions, particularly bulk density (since bulk density data are used to calculate carbon stocks) of tilled croplands had returned to their original pre-tillage state 45 Deep sampling to a depth of 90 cm with a sampling depth interval of 30 cm was used for the study as previous studies in the area were limited to only 0-30 cm depth. In addition, the deep sampling intervals were assumed to be consistent with the current standard soil depth of 30 cm as proposed for C accounting studies 47,48 . Systematic sampling was adopted for soil sample collection with transects established in each land use type. Soil samples were collected from three representative plots (50 × 50 m) on the established transect for each land use type. In all land use types, undisturbed samples were first collected using open-faced coring tube, before auger sampling from the same point. A hand-pushed auger was used in collecting the auger samples. Replicate plots within each land use type were approximately 400 m apart and the experimental plots were of the same lithology and management. Within each plot (replicate) in a land use type, auger samples were collected at each depth from four points of a soil profile pit (1.5 × 1 m), along the already established transect, giving four sampling positions  Bulk density determination was carried out using the core samples. Bulk density samples were collected with the aid of core samplers, starting from the lowest soil depth (60-90 cm) to the topmost soil depth (0-30 cm). The auger samples were first air-dried, followed by manually removing the visible roots, twigs, debris and leaves, and finally sieving the soil using a 2.0 mm mesh screen. The < 2 mm samples were then subjected to further laboratory analysis.
Laboratory analysis. Soil organic carbon content was determined using modified Walkley and Black wet oxidation method with H 2 SO 4 -K 2 Cr 2 O 7 followed by residual titration with 1 N HCl 49 . Total nitrogen was determined by the modified macro Kjeldahl digestion method 50 . Soil pH was measured in soil-water (1:2.5) suspensions 51 . Cation exchange capacity (CEC) determination was by NH 4 OAC (pH 7) displacement method 52 . Bulk density was analyzed using core method 53 . Particle size distribution was determined by the hydrometer method 54 using sodium hexametaphosphate as a dispersant. All measurements were taken in triplicates for improved accuracy.
Calculations, estimations and statistical analyses. The SOC and TN stock in each land use type was calculated with the formula: where concentration (%) is the percentage concentration of carbon or nitrogen. The total SOC and TN stocks to the depth of 90 cm across locations in each land type use was calculated by the summation of SOC and TN stocks in the 0-30, 30-60, and 60-90 cm soil depths 55 .
In Geregera and Haikihelet locations, using the grazing land as a baseline, SOC and TN stocks accumulation in exclosure within the same soil depth were obtained by calculating the difference in SOC and TN between exclosure and grazing land. Due to variation in periods of the different land use types, rate of SOC and TN  www.nature.com/scientificreports/ stocks accumulation for each soil depth of the exclosure land use type was calculated by dividing the estimated accumulation values by the presumed period of exclosure establishment 56 . The average age duration of Geregera and Haikihelet exclosures were given as 10 and 6 years respectively 42 . This simply implies the age duration in years since the exclosures were established from degraded grazing land. This information was based on the oral feedback from farmers (aged 60 years and above) in the study locations.
In Desa'a and Hugumburda locations, SOC and TN losses due to deforestation were estimated by subtracting the total SOC and TN stocks in forestland from its equivalent depth in grazing land or cropland. Thereafter, the calculated loss values were divided by the presumed period of years following land use conversion to get SOC and TN losses per year. The CO 2 emission as a result of forest conversion to grazing land and cropland was then established on the basis of the underlying SOC and CO 2 relationship as stated by 57 ; which states that 1 Mg ha −1 increase in soil carbon signifies removal of 3.67 Mg of CO 2 from the atmosphere. For the purpose of this study, we are focusing only on C lost as CO 2 emission without consideration the C losses through erosion, and leaching in the form of dissolved organic C and sediment accumulation. From the obtained results of SOC and TN concentrations and stocks, the distribution trend was explained in the form of high, intermediate and low across the different land use types in all study locations for ease of comparison.
Considering that soil carbon quantity is specifically quantified in a particular soil depth for the purpose of C accounting and budgeting 47 , effects of land use on SOC and TN concentration and stock, CEC, pH, bulk density, was investigated, comparing them across same depth within site based on two-way analysis of variance (ANOVA). Log transformation of data was carried out before ANOVA whenever assumptions of normality and homogeneity of variances within a group were not obtained. Significant differences (p ≤ 0.05) were determined using Tukey's honest significant difference (Tukey's HSD) post hoc test. All the tests were carried using STATIS-TICA (Version 12.0, StatSoft GmbH, Hamburg, Germany). Factor analysis was performed using version 2014 of XLSTAT (Addinsoft, Paris, France).

Results
Soil organic carbon and total nitrogen (stocks and contents), and C:N ratios in top versus subsoils. Soil organic carbon and total nitrogen stocks and concentrations displayed a similar pattern-decreasing with soil depth among the land use types in all study locations (Fig. 3, Table 2). In Desa'a and Hugumburda, the C stocks per land use type is ranked as, forest (122.98 and 39.26 Mg C ha −1 ) > grazing land (72. a' a" a' a"   (Fig. 3A). The TN stock showed significant (p ≤ 0.05) difference between land use types across locations, following a similar distribution pattern with SOC stock (Fig. 3B). Across all locations, SOC and TN stock decreased with soil depth, with high values in forest lands, medium in grazing lands and exclosures, and low in crop lands. The SOC and TN content (%) followed similar trend as the SOC and TN stock distribution (Table 2), decreasing with soil depth in all land use types across locations. There was no clear distribution pattern of C:N ratio with depth under different land use types. The soil C:N ratios were high in forest and low in cropland across locations, except for the cropland in Desa'a which recorded higher C:N ratio than forest (Table 2). Across all locations, the soil C:N ratios ranged between 22.80 and 7.04.

Accumulation and loss of SOC and TN stock.
With exclosure establishment, high SOC stock accumulation in Geregera (16.57 Mg C ha −1 ) and Haikihelet (64.20 Mg C ha −1 ) was observed ( Table 3). The high SOC accumulation also accounted for a high SOC accumulation rate of 6.88 Mg C ha −1 yr −1 in Geregera and 10.7 Mg C ha −1 in Haikihelet. The TN accumulation and rate of TN accumulation for Geregera and Haikihelet were 0.18 Mg N ha −1 and 0.02 Mg TN ha −1 yr −1 and 1.29 Mg N ha −1 and 0.22 Mg TN ha −1 yr −1 respec- Table 2. Soil organic carbon and total nitrogen concentrations, C:N ratio, cation exchange capacity and pH under different land use types and soil depths in the study locations. ± Mean followed by standard errors. Letters after the standard errors indicate significant differences (P < 0.05) between land uses at 0-30 cm (a), 30-60 cm (a′) and 60-90 cm (a″). In Desa'a and Hugumburda, conversion of forest to grazing land and cropland accounted for huge SOC stock depletion amounting to 30 to 50% of the SOC in the topsoil layer. Forest conversion to grazing land in Desa'a resulted to a reduction in total SOC and TN stock of 3.50 Mg C ha −1 yr −1 and 0.36 Mg N ha −1 yr −1 in grazing land, and 9.04 Mg C ha −1 yr −1 and 0.99 Mg N ha −1 yr −1 in cropland respectively. Hugumburda followed the same pattern of SOC and TN stock loss, though with less magnitude accounting for estimated total SOC and TN stock loss of 0.61 Mg C ha −1 yr −1 and 0.03 Mg N ha −1 yr −1 in grazing land, and 2.05 Mg C ha −1 yr −1 and 0.12 Mg N ha −1 yr −1 in cropland respectively (Table 4).  Table 2).
In Vertisols of Desa'a location, no land use effect on CEC was observed, whereas in all other locations, predominated by Cambisols, croplands displayed higher CEC values more than grazing land, with the exception of Geregera. There was an observed decrease in CEC content across soil depths in all land use types in studied locations. www.nature.com/scientificreports/ Land use types and soil depth affected the pH value of the soils, though with narrow margin across locations. The highest soil pH of 8.10 was recorded in Geregera cropland 0-30 and Haikihelet cropland 60-90 while the least soil pH value of 7.2 was recorded in subsoil (30-60 cm) of Desa'a grazing land (Table 2). Generally, the soil pH was in the slightly alkaline range in the study area (Table 2).
Particle size distribution of the soils showed significant (p ≤ 0.05) variations across depths under different land use types (Table 5) with an exception in Hugumburda where total sand and silt fractions showed no significant difference (p ≥ 0.05) across depths among land use types. Predominance of total sand fraction in Cambisols of Hugumburda and Geregera in different land use types was observed. In Cambisols of Haikihelet, total sand fraction was dominant in exclosure, while clay and silt fractions were dominant in grazing land and cropland respectively. In Vertisols of Desa'a, silt fraction was dominant in forestland while clay fraction was dominant in grazing land and cropland. Clay fraction increased with depth in most land use types across locations except in Hugumburda (forest and grazing land), Haikihelet (forest, grazing land and cropland) and Desa'a (forest) ( Table 5).
Bulk density (BD) differed significantly (p ≤ 0.05) with depths across different land use types in the studied locations. Similarity existed between topsoil BD of grazing land and cropland. Across locations, the highest BD value of 1.53 Mg/m 3 was recorded in 60-90 cm of Haikihelet exclosure while the lowest BD value of 1.06 Mg/ Table 5. Soil physical properties under different land uses and soil depths in the study locations. ± Mean followed by standard errors. Letters after the standard errors indicate significant differences (p < 0.05) between land uses at 0-30 cm (a), 30-60 cm (a′) and 60-90 cm (a″). www.nature.com/scientificreports/ m 3 was recorded in both 0-30 and 30-60 cm depth in Desa'a forestland. Significant increase in BD with depth was observed across locations (Table 5).

Land use Depth (cm) Total sand (%) Silt (%) Clay (%)
Factor analysis across locations. In Geregera, the first factor axis (F1) of the biplots relates to plots gradient from grazing land to cropland, however, there is similarity between the plots of grazing land and forest land. Most of the studied soil properties (SOC, TN, C:N ratio, CEC, SOC and TN stock, pH, % silt and % clay) were higher in exclosure and grazing land compared to cropland (Fig. 4A). Notably, cropland soils are characterized by loam, sandy loam, silty clay loam and sandy clay loam texture for Hugumburda, Haikihelet, Desa'a and Geregera respectively (Table 2), in addition to low SOC content as observed in the second factor axis (F2) which explained 23.80% of the total variance. In general, this relates to a particular gradient of low SOC (cropland plots) to high SOC (exclosure plots) (Fig. 4A). In Desa'a, the first factor axis (F1) of the biplots followed particular pattern modulated by percent clay distribution. Cropland has higher percent clay content, followed by grazing land, with forest land recording the least percent clay content (Fig. 4B). The second factor axis (F2) of the biplots indicates a pattern of inclination from forest to cropland plots. However, a clear pattern was observed, starting from forest with high SOC and high sand fraction to cropland with low SOC and low sand fraction (Fig. 4B).
The biplots in Haikihelet indicated that the first factor axis (F1) exhibited a clear pattern of inclination from exclosure to cropland plots. The second factor axis (F2) followed a different pattern of increment from cropland to grazing land, corresponding to a gradient of low to high SOC pool. The grazing land is characterized by high SOC content, SOC stock and high clay content. (Fig. 4C). www.nature.com/scientificreports/ It was observed that the biplots in Hugumburda both for the first factor axis (F1) and second factor axis (F2) of the biplots explained approximately 65% of the variances in the components (Fig. 5A), thus necessecitating to incorporate third factor axis (F3) (Fig. 5B). The first factor axis (F1) explained 41.25% of the total variance in forest and partly in grazing land, consisiting of variables indicative of soil nutrient availability (SOC, TN, SOC stock, TN stock, CEC, and C:N ratio) (Fig. 5A). The second factor axis (F2) which explained 23.62% of the total variance, was characterized by % clay, BD and pH in cropland and grazing land. The third factor axis (F3) explained 19.70% of total variation, and was characterized by pH and BD in grazing land and cropland and so, reflects the impact of anthropogenic activities (Fig. 5B).

Discussion
Significant difference in SOC and TN stocks was observed among various land use types across depths, with clear differences in distribution trend across locations. The high SOC and TN concentration recorded in topsoil (0-30 cm) of forest and exclosure could be as a result of minimal disturbance in these ecosystems, litter accumulation from trees and shrubs, below ground litter and high biomass cover [58][59][60] . In addition, SOC and TN distribution at lower depth in most land use types were usually low at 60-90 cm depth, thus indicating that effect of land-use was mainly limited to the upper soil layers. This did not support our first hypothesis which states that the impact of land use is not limited to the topsoil. The significantly lower concentrations of SOC and TN especially in cropland soils across depths could be attributed to unsustainable farm practices like total harvesting without residue retention thus exposing the soil to incidences of soil and water erosion, residue burning and intensive tillage operations which exacerbates decomposition and high rate of SOM oxidation due to continuous cultivation [61][62][63] . However, with good management practices in croplands, C retention and stabilization can be enhanced.
A possible attribution of high SOC contents at Haikihelet and Desa'a is the high clay content recorded in these locations (See Table 5). In this study, both SOC and clay contents were found to be higher in Vertisols (clay dominated) than Cambisols (sandy loam dominated). This finding is in line with works of 64 and 65 who reported that clay textured soils had higher SOC content in studies assessing distribution of SOC levels and structural indices under contrasting land use types in southeastern Nigeria.
The forest sequestered more SOC stock than other studied land use systems, with high sequestration in topsoils. This is in line with the report of 16 that forest soils are great C pools of terrestrial ecosystems in the global C cycle. This implies a high risk of CO 2 release from these forest topsoils if they are eventually deforested or converted to cropland. The low SOC stocks in the cropland are attributable to the total harvest, tillage activities in addition to leaching and erosion losses and reduced organic material going back to the soil, soil and water erosion leading to loss of SOM, regular tillage and cropping activities accounting for high oxidation rates of SOM, burning of crop residues 19,61,66 .
Our results show that C:N ratio was affected by land use types, but there was no definite distribution trend. This suggests that the sole use of C:N ratio as a SOM quality indicator is limited and quite misleading 67 . The lowest C:N ratio in 30-60 cm depth in cropland soil of Hugumburda (7.66) corresponds to a very low SOC content of 0.25% ( Table 2). The ratio was much narrower in croplands (with the exception of Desa'a cropland) than other land use types, which is an indication of high mineralization and oxidation rate in cropland soils. Decline in C:N ratio with soil depth is evident in most agricultural soils 68 . Interpretation of changes in C:N ratio due to in land use changes or management practices is complex and has been suggested to be better treated separately from SOC and TN concentrations and stocks, and on a case by case basis for clear understanding 67 . www.nature.com/scientificreports/ At Geregera and Haikihelet, taking grazing land as the baseline, increase in SOC and TN stock accumulation as a result of exclosure establishment was recorded. The observed improvement in SOC and TN stock in 6-and 10-year-old exclosure in Haikihelet and Geregera respectively is attributed to increase in organic inputs due to vegetation restoration and restriction of animal grazing on exclosures. This firmly supports section of our third hypothesis which states that exclosure establishment on degraded ecosystems results in SOC restoration. The implication of our result is that long age duration of exclosures may not necessarily result to remarkable replenishment of soil nutrients on previously degraded grazing land in compared to short-term exclosures. Sitespecific characteristics and micro-climatic conditions across locations might have contributed to these variations in SOC accumulation rates.
Most of the SOC and TN stocks losses were in the 0-30 cm topsoil layer across land use types (Table 4). Similar trend in SOC loss has been previously reported by 34 who indicated that forest conversion to crop land, open grazing, and plantation accounted for an estimated decline in SOC stock in the topsoil layer amounting to 0-63% in cropland, 0-23% in open grazing land, and 17-83% in plantation. This confirms a section of our third hypothesis which states that SOC concentrations and stocks decrease after conversion of natural forests to cultivated lands. In the dry Afromontane remnant pristine forests in northwest Ethiopia, huge reduction in SOC stock of up to 87% and 50% with the conversion of forest to cropland at Katassi and Gelawdios sites respectively was reported by 59 . This portends a huge threat to global warming in the face of climate change.
Overall, with forest conversion to cropland and grazing land, the estimated CO 2 emission as obtained in this study is huge and capable of contributing to atmospheric greenhouse gas effect. The CO 2 emissions decreased with soil depth with higher emissions in cropland compared to grazing land soils. Notably, CO 2 fluxes decreases appreciably with depth though not significantly contributing to surface fluxes 69,70 . This implies that any form of subsoil disturbance could affect the deep subsoil CO 2 reservoir. Thus, mobility of CO 2 in the subsoil to the surface soil is impaired and may be entrapped in soil pores and solution if undisturbed, or used by subsoil autotrophs 19 . In cropland soils with high CO 2 emissions, these CO 2 loss effects can be compensated by the accrual of deep root C inputs from deep-rooting crops. Recent studies of deep-rooted perennial grasses planted in C-poor soils reported no effect of these crops on surface CO 2 fluxes in different soil types 71,72 . Nevertheless, the value of SOC loss in our study indicates that loss of SOC may not only be as a result of CO 2 emission to the atmosphere 73 but can as well be lost due to leaching in the form of dissolved organic carbon, erosion and sediment accumulation, which were not considered in this study. Thus, the actual fate of this SOC loss across landscape in semi-arid area of northern Ethiopia is still not well-known and there is need for further detailed investigation.
Rainfall has been reported as the main governing factor of SOM and TN content distribution in Sub-Saharan tropical soils of East Africa 74 . This was affirmed by our result of overall mean high SOC and TN stock especially in Desa'a with high rainfall and low temperature compared to other locations (See Figs. 6 and 7). Observably, SOC stock increased with increasing mean annual precipitation (MAP) and decreased appreciably with increasing mean annual temperature (MAT) (See Figs. 6 and 7). Various authors have reported positive correlation of SOC stock with MAP but with negative correlation with MAT 59,75 . Assefa et al. 59 reported high SOC stock values in www.nature.com/scientificreports/ areas with high MAP compared to areas with low MAP in northwest Ethiopia. Estimated SOM content ranging between 0.5-3.0 and 10-13% for tropical soils of Sub-Saharan Africa (high temperature) and temperate soils of Europe/America (low temperature) respectively has been reported 2 . Global distribution of SOC stock follows a pattern, increasing from temperate (cooler) regions to tropical and sub-tropical (hotter) regions 76 . Temperature and precipitation remain the two major environmental factors affecting SOC concentration and stock within the complexity of land use change (LUC)-SOC distribution nexus 77,78 . In our study, MAP has more impact in terms of modulating SOC stock compared to MAT (See Fig. 6). This partly supports our second hypothesis that climate and land use history will serve as possible indicators to modulate the amount of change in SOC stocks. Another remarkable outlook in this study in terms of drivers of SOC distribution was provided by the clay fraction data on basis of occurrence or proportion. Thus, soils with high clay content recorded high SOC concentration and stock compared to soils with low clay content. This finding is in agreement with the work of 67 in a study to assess the C:N ratios following land use change in Brazil.
In Cambisols of Hugumburda, Haikihelet and Geregera, the overall mean total sand fraction was high in natural (forest) and semi-natural (exclosure) ecosystems, with high proportions in sub-soil layers in most land use types excluding cropland at Geregera. This is in contrast with the findings of 79 who reported high sand fraction in grazing land, followed by agroforestry and cropland in Nitosols of southern Ethiopia. In Vertisols of Desa'a, high clay fraction was observed in all the land use types with appreciable increase with depth. Differences in soil types and micro-climatic conditions might be responsible for these variations. Bockheim 80 , Ukaegbu et al. 81 and Okolo et al. 82 reported that soils formed on the same parent material within an ecological region are complexly linked to landscape and thus display substantial variations in soil properties. Furthermore, differences in BD was observed across locations in different land use types. Low BD in natural (forests) and semi-natural (exclosures) ecosystems, could be attributed to constant input of high soil organic residues on the upper layer of the soil [83][84][85][86] . The contribution of tree roots to the subsoil organic matter (OM) accumulation, including root litter decomposition leads to the decrease in BD in forests [87][88][89] .

Conclusions
The total SOC and TN concentrations and stocks were high in natural forest, intermediate in exclosure and grazing land, and low in croplands, and generally decreased with increasing depth in allland use types. Across soil depths and land use types, SOC and TN sequestration was higher in Cambisols than Vertisols, with clay content and MAP rather than C:N ratio alone being the most meaningful indices for SOC storage and soil quality assessment. Conversion of forest to cropland resulted to significant losses of SOC and TN with considerable amount of CO 2 emission which contributes to change in climate. Exclosure establishment supported restoration  www.nature.com/scientificreports/ of degraded grazing lands with recovery of SOC and TN stocks especially in the topsoil layer (Fig. 7). Thus, exclosure establishment could be a sustainable way to reverse soil fertility decline due to its C and N sequestration potentials. Additionally, more attention needs to be placed not only on the amount of SOC sequestration potential under different ecosystems and land use types in semi-arid area of northern Ethiopia, but also to ensure that they remain undisturbed for long periods of time, with mechanisms to detect differences before commencement of carbon trading schemes.

Data availability
The datasets used for this study are available from the corresponding author on reasonable request.